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Principles of
ANGIOSPERM
TAXONOMY
P. H, DAVIS, PH.D., D.Se.
University oj Edinburgh
V. H. HEYWOOD, PH.D., D.Se.
University of Liverpool
1991
KRI EGER PUBLISHING COMPANY
Malabar, Florida, USA
Exclusive North American Distributor
Chapter 4
THE CONCEPT OF CHARACTERS
"A knowledge of the relative importance of characters can only
be acquired by lcng study; and without a due appreciation of
their value no natural group can be defined."
J. D. HOOKER, . 1855
INTRODUCTION
Taxonomy, in so far as it is concerned with variation in organisms and their
classification, deals with characters. Characters provide the taxonomic
evidence for natural classification in the Adansonian sense and a large part
of the evidence for evolutionary ("phylogenetic") classification. As Cain has
recently stressed, the basis for a natural classification is entirely the characters
available. * It is therefore surprising that the problems of characters as such
should have received so little consideration by taxonomists, since the success
of most taxonomic studies must depend to a very large extent on what
attributes are selected and considered as characters, how they are tested and
treated, and whether they are relevant for the purpose in hand. It should be
pointed out straightaway that the concept of characters as recognisable,
separate entities is a product of man's necessity to communicate and there:
fore to describe. This explains why characters are so difficult to define. This
aspect is returned to later.
Although it is organisms (or life cycles of organisms) which are classjfied, t
it is their characters which provide the evidence used in classification.
Ideally the whole organism (i.e. all its attributes) should be employed, but
since each individual possesses thousands of potential characters, practical
limitations impose a restriction on the numbers used. As Muyr the
number is limited by the patience of the investigator. This leads on to the
problems of selection. Lam (1959) believes that it is probably not possible to
make a fully relevant choice if ultimate. goal is a completely natural
classification!' Moreover, different selections of attributes will produce
Throughout this chapter, natural classification is understood as meaning
phenetic.
t More strictly speaking, it is our knowledge of them at. any time that is
classified.
INTRODUCTION III
different classifkations or different versions of the "same" classification. The
selection of features made will, in turn, depend on a number of factors such as
the purpose of the classification (an aspect often ignored. cf. Crawshay
Williams, 1961), the general convenience of the scientist, the availability of
characters, and traditional usage.
Most classification hitherto has employed morphological and anatomical
characters as its major source of evidence, but as is shown later in this
chapter other kinds of features are now being increasingly empl0yed, and it is
by no means certain that the results will be acceptable for general purposes.
It has, for example, so far been tacitly assumed that natural classifications
will be morphologically expressible and comprise non-overlapping groups.
Both these assumptions are open to question.
All classification, as Lam has put it, must be a compromise because of the
particular selection of characters and the degree of precision and unity with
which they can be defined. Human limitations are in fact the main limiting
factor.
For general purposes it is widely, although by no means universally,
agreed that a maximum-attribute classification is the most useful: it is the
classification which allows the maximum number of generalisations to be
made from it (Gilmour & Turrill, 1941). As Bremekamp (1939) has explained,
it is the most serviceable because it conveys the maximum possible amou!}t of
information about the groups it contains.
Such classifications are, in theory at least, undertaken following the
precepts established by Michel Adanson (p. 23) and are frequently called
Adansonian. The two essential postulates are: (1) in constructing the classi
fication each attribute selected is given equal weight; (2) taxa are based on
correlations between these attributes. Further, the taxonomic groups derived
by these means are of the class of concepts which have been termed polythetic
(Sneath, 1962) "in which no single attribute is in theory sufficient and necessary
for membership in the group so long as the members share ahigh proportion
of characters" (Snc.ath & Sokal, 1962).
Many taxonomists find great difficuJty in accepting these premises,
largely because they regard them as and therefore un
acceptable in this day and age; but failure to accept them is to distort classi
fication by the introduction of phyletic, logical or other sorts of weighting
principles. Such principles may be justified or are indeed necessary for the
construction of special classifications (e.g. phyletic), but are not compatible
with the aims of a general classification. In other words, they serve a different
purpose. By giving greater weight to particular evidence, a classification
ceases to be a general one.
These Adansonian principles for the construction of phenetic classifica
tions have received great support from the almost spectacular development
in the past five or six years of what Sneath and Sokal have called "Numerical
Taxonomy"-"the numerical evaluation of the affinity or similarity between
112 THE CONCEPT OF CHARACTERS [Ch.4
taxonomic units and the ordering of these units into taxa on the basis of their
affitiities"-which is essentially an extension of Adansonian classification
using mathematical procedures with the primary aims of repeatability and
objectivity: The methods and applicability of numerical taxonomy are dis
cussed in a later section (p. 131).
The groups recognised in natural classifications are obviously based on
overall similarity. Mathematical tecluiiques, often employing computers, are
now being used for making the character correlations (see p. 135). The
groups so delimited are not, however, to be interpreted as necessarily phyletic,
due to a number of factors such as convergence, functional correlation and
difficulties of ascertaining homologies. In considering the taxonomic . value
of the kinds of characters which are discussed in this chapter, a clear dis
tinction must be made between their role in maximum-attribute (natural)
classifications and special (phylogenetic) classifications (phenetic and phyletic
respectively, as A. J. Cain (1959d) has recently termed them). The conversion
of a phenetic classification into a phyletic one involves a further series of
problems of interpreting characters and weighting them (see p. 127). It is
important not to mix the two approaches, as has frequently been done in
the past. The separation of phenetic and phylogenetic classifications is not
generally accepted today due to the intense conviction that classification
should be evolutionary and that it is the only sOft of classification which
should occupy the serious attention of scientific taxonomists. We must
insist, however, that it is not inconsistent with evolutionary thinking to
encourage this separation, and advocate great concentration on the primary
task of producing general classifications based on maximum co-variation of
attributes/or general purposes. In flowering plants the possibilities ofachieving
a phyletic arrangement or fully evolutionary classification as it is normally
understood today are limited for reasons which are discussed below. This is
not to deny, however, that one of the major activities of taxonomy is the
evolutionary interpretation of evidence, and part of this chapter is devoted to
considering how this may be done.
DEFINITION OF CHARACTERS
From an evolutionary point of view characters should probably not be
considered in isolation but in combination, since it is on character-combina
tions that natural selection operates. This is particularly true if one is con
sidering the possible selective or functional value of characters, but for
general taxonomic purposes it is necessary to deal with unit characters which
can then be treated quantitatively. As is explained elsewhere (Chapter 2),
the taxonomist's assessment of similarity is made to a large extent intuitively,
and only those characters which appear to be differential between organisms
or groups of them are set down on paper. The current tendency in taxonomy
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DEFINITION OF CHARACTERS ll3
is to analyse these intuitive processes and rationalise them by distinguishing
the individual characters which, the taxonomist assesses subconsdously
together. TlUs raises the problem of the definition of unit characters. It
seems unlikely, however, that strict or objective definition can be made, due
to the difficulties of deciding in many cases whether any particular attribute
. of a plant should be regarded as one or more for the purposes of
making comparisons. Many of the difficulties arise from a lack of knowledge
of the anatomy and morphology of the various organs and features of plants.
A general definition of a character is: any attribute (or descriptive phrase)
referring to form, structure or behaviour whiCh the taxonomist separates from
the whole organism for a particular purpose such as comparison or interpretation.
Cain & Harrison (1958) describe a single character as "anything that can
be considered as a 'variable independent of any other thing considered at the
same time".
Characters as such are strictly speaking abstract entities: it is their
expressions or states that taxonomists deal with. "Sepal length" for instance
may be considered as a character; but "sepal length 12mm," is an expression
of that character. For purposes of comparison, a character must be divisible
into two or more expressions or states (including here presence or absence
in qualitative features).
For practical purposes a character may be defined as any feature whose
expression can be measured, counted or otherwjse assessed (cf. Hedberg,
1957). Examples are: seed wdgfit, ratIo of corolla to calyx length, number of
teeth on a leaf, presence or absence of tubercles . in leafaxils, number of
transverse septa in anthers, to <;ite a few at random. One of the features of
modern taxonomy is the wide range of characters that are being employed.
The question "what is a character?" is, however, more difficult to answer
than the above definitions suggest. Many characters may be regarded as
compound or multiple. For the purposes of numerical taxonomy Sneath &
Sokal (1962) state that characters should be unit characters or if they are
multiple they should be broken down into unit characters. As a working
definition of a unit character they propose "a taxonomic character of two
or more states, which within the study at hand cannot be further subdivided
logically (except for subdivision brought about by changes in the method of
coding the states)". However theoretically sound this may be, there are
bound to be acute difficulties in making a decision about the logic of
up characters into units, a limiting factor being our own state of knowledge,
skin and perception. This is one of the most' serious weaknesses of any
attempts to break down organisms into characters for the purposes of
This method is essentially similar to what Matthews (1962) has recently
termed neural science-the private, individual knowledge each man obtains for
himself through his senses. It is contrasted with formal science-the knowledge
obtained by measurement, testing experiment, and the particular kind .of logic
based on these evidences of reality. .
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114 THE CONCEPT OF CHARACTERS [Ch.4
objective comparisons. In other words, although the comparison of characters
may be objective, the selection of what is to be regarded as a character must
necessarily be largely subjective.
Let us consider an actual example: how many characters are involved in a
leaf shape, for instance? The expression "obovate" may be controlled by
distinct genes, some of which influence length, others of which influence
width, and all of which probably interact. The concept "obovate" may be
divided into components dealing with length, maximum breadth, position of
maximum breadth, breadth at various intervals, length/breadth ratio, etc.
the shape being the product of differential growth rates, themselves controlle4
by genetically determined hormone systems. In many cases it may be
sufficient to take the general outline, i.e. obovate, as a single character;
in other cases it may have to be broken down and analysed biometrically.
Leaf dentation and the form of the apex may be treated as different characters
since they may vary independently of general outline. In compound leaves
the problems are correspondingly complicated.
It follows from what we have just said that no precise general answer
can be given to the question "what is a character?" This can only be considered
in individual cases and what we treat as a character will depend on what we
want to use it for. Even apparent absence of differential characters or expres
sions between individuals or groups need not indicate that they are identical:
differences may well come to light after detailed study. It has been remarked
that one will always find characters for separation if one tries hard eno\lgh
and, one might add, find that characters used for separation do not hold when
more material is examined!
Characters may be considered from two viewpoints corresponding to two
main activities of taxonomy:
(a) Identification, characterisation and delimitation of species.
(6) Grouping (classification) of these species into higher taxa (natural
groups), i.e. into a system that as far as possible expresses their natural
relationships.
Corresponding with these activities, characters may be regarded as
analytic or synthetic: in each procedure the actual characters used are
usually different in each, particular instance, although there is nothing
inherently different in them; it is rather that different kinds of character are
selected. In identification and characterisation the taxonomist seeks to employ
diagnostic or key characters, i.e. those of limited ou.:urrence, selected so that
their use alQl1e is sufficient for reaching a correct diagnosis. In arriving at a
decision about the delimitation of species numerous characters may be used;
many of them of a highly specialised nature which do not lend themselves to
easy description or recognition, e.g. chemical, cytological, physiological
characters or those which show only a statistically significant difference.
These characters are not normally given in descriptions (in Floras, revisions,
etc.) which often have to be short and precise. Descriptions are really
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DEFINITION OF CHARACTERS 115
abstracts for practical convenience-they give some, at least, of the characters
which are easily recognisable and comparable, not all those features which
may have been used in making a decision about status. In this activity,
taxonomists not only gather information (i.e. characters) together,but
present them predigested. This is often not realised, especially by non
taxonomists who tend to assume that a particular classification stands or
falls by the evidence quoted. It does raise the problem, however, of the amount
of data that should be presented in any ,articular context, and clearly the
purpose of the publication is a major consideration.
The purpose of a Flora is still today primarily to indicate what plants
occur in a particular area, how they may be identified (by keys and descrip
tions) and what scientific names refer to them (correct nomenclature and
synonymy). Certain subsidiary information is normally included, such as
ecological and distributional data and chromosome counts (cf. discussion in
Heywood, 1960a: 153-154). Diagnostic characters make the process of
identification in such Floras easier. Discussion about them and the other
characters which mayor may not be given in the description is out of place
and hinders rather than hastens the purposes of identification. Such informa
tion may be published elsewhere. as it often is in detailed monographic
studies.
1. Analytic and Synthetic Characters
As noted above, characters used in identification, characterisation and
delimitation may be termed analytic, following Just (1946). In the classifi
cation of the results of these activities, ,synthetic characters are used, i.e.
characters of a constant nature which unlike diagnostic characters are of wide
The analytic or diagnostic characters which serve to recognise a
group are seldom of use in synthesising the group along with others into a
higher group. Fpr this purpose one needs'characters whose constancy of
occurrence increases the higher the position of the group in the hierarchy.
Those characters used to synthesise lower groups into higher groups serve at
the same time to distinguish the higher groups from others of the same iank.
This was recognised by Darwin who noted that the classificatory value
of a character which has a wide occurrence (in many forms), as opposed
to one which has restricted occurrence, was in terms of hierarchical value.
In other words, characters helped to determine different ranks. Characters
which serve to separate genera in one group are of no value for such a purpose
in other groups (cf. Darwin, Origin, Ch. XllI, discussion of Robert Brown
and characters of the Connaraceae).
As already said, there is no inheren t di fTerence between analytic and syn
thetic characters: the difference is more one of the particular use made of
them in a particular case. Charactcrs which differ belween taxa are usually
similar in kind to those that arc found varying within species populations
as Simpson (l953a) has noted. He comments that "one needs only to take the
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116 THE CONCEPT OF CHARACTERS [Ch.4
diagnosis of virtually any taxonomic group to find that the same diagnostic
characters are variable in other and related taxonomic of the same
or lower hierarchic rank and that eventually they can be traced down to
variation in populations" (1953a:' 1(0).
On the other hand, certain features which characterise higher groupings in
the Angiosperms, such as. superior versus inferior ovary, are seldom today
found to vary within populations. These are the so-called "constitutive"
characters discussed on p; 122.
The distinction between analysis and synthesis in taxonomy is not always
appreciated. Genera, for example, are syntheses of species, but having
inherited most of our genera from previous generations of taxonomists, we
find that they have, over the years, been enlarged by accretion and not by
deliberate resynthesis from species. When 'a periodical review of genera is
undertaken, this resynthesis often leads to an alteration in generic limits and
consequent change in status. As an example of the use of analysis when
. synthesis was required, we may cite Williams's revision of Arenaria in which
he divided the greater part of the genus into 3 subgenera based on supposed
differences in structure of staminal glands, thereby disregarding the
ledged affinities of the species as judged by overall resemblances. This is an
example of analysis by a weighting of characters and contrasts with the
synthetic treatment of the same group by Fenzl (1840) and McNeill (1962). It
should be pointed out that staminal gland structure serves as a useful diag
nostic (analytic) character to separate species Arenaria armeniaca and
A. zargariana) within the genus after it has been revised by synthetic pro
cedures.
2. Qualitative ,and Quantitative Characters
A distinction is often made between qualitative (including presence or
absence) characters and quantitative characters for descriptive purposes.-Thus
features assessed by size, length, etc. are quantitative while those relating
to form, for example, can be considered as qualitative. Such characters as
opposite/alternate leaves and placentation types are clearly qualitative.
Often, however, the distinction is more apparent than real: many qualitative
characters may also be expressed quantitatively (cf. leaf shape discussed
above), and it is perhaps more useful to consider characters according to
whether they may be assessed directly or not (Heslop-Harrisop, 1952b).
Characters may be directly by counting (by number), by simp1e
measurements of line<rr dimensions (by size), or by angles (by shape). These
are the most satisfactory from a practical point of view in that they can be'
easily scored, described and processed. The advantages of such characters
. for the purposes of biometrical or statistical comparison are obvious. (The
variation of characters in populations and its causes are considered in Chapter
11).
(a) Number. Different numbers or ranges of numbers of parts (such as
117 DEFINITION OF CHARACTEBS
leaves, flower parts) are COmIilonly employed in taxonomic descriptions.
The term meristic variation is applied to such cases where the variate can only
assume discrete values. Seed number when of low value is assessed in this
way too, but when the values are high, assessment by weight of numerical
units (e.g. 100 seeds) is often employed. By their very nature the recording
of such numerical characters seldom presents particular problems.
(b) Size. Linear dimensions, too, are commonly given in descriptions,
often with a conventional indication of range and extremes, e.g. petals
(3-) 5-7 (-9) mm. Overlapping values between two taa in such characters
(especially vegetative) are frequent and they are not the most
useful for taxonomic separation. Apparent differences may be due to in
adequate sampling, especially when dealing with conventional herbarium
collections. On the other hand, the differences may be on quite a small
scale (such as anther length in Centaurium spp.) and afford a reliable
criteriqn. Such small-scale differences are frequently neglected since they are
not evident without precise measurement under strong magnification. Then
there is the psychological problem, too, that small-scale characters tend to
be considered unimportant, while immediately obvious ones are regarded as
valuable. On the other hand, many botanists have considered characters of
primary importance just because they are difficult to observe] Neither claim
is valid-the importance of a character is to be judged by other criteria, as is
discussed below. It is also possible that above certain dimensions differences
between organs may be ignored, particularly when other species in the genus
have relatively small organs.
In many instances differences in linear dimensions (between popUlations)
may only be statistically significant and consequently of little value as
primary recognition characters.
(c) Shape. As"we have already seen, in shape present consider
able problems although they are often more valuable for diagnostic purposeS
than size differences. They often contribute more to differences in facies
and are instrumental in causing taxa to be recognised as distinct, but their
description is correspondingly difficult. The techniques involved in the detailed
specification of leaf shape by numerical indices are discussed later, but some
more general problems have to be considered at this point. Simple leaf
shapes, such as ovate, lanceolate, linear, etc., are bedevilled by problems of
terminology arid different usages. It soon, becomes evident, even to relatively
inexperienced taxonomists, that there is a lack of precision in the use of the '
various leaf-shape terms and inconsistency of usage between different
Floras, etc. Nothing, of course, can be done to alter past usage although it is
often possible to learn what conventions have been employed in particular
countries or by particular authors. It is strongly to be recommended that
today precise systems for designating conventional shapes should be followed,
such as those given by Stearn (1956). There are signs that the matter is
receiving international consideration. (See p. 271).
118 THE CONCEPT OF CHARACTERS [Ch.4
Many characters may not be susceptible to direct numerical assessment,
such as colour, texture, indumentum, smell; taste, patterning, etc. They are
none the less widely used in taxonomy, as is in the next and
clearly precise means of describing them are necessary.
Colour can be treated in several ways: the use of international standards
such as the Ridgway and Royal Horticultural Sociery colour charts, whereby
colour is divided into a number of arbitrary named classes and specimens are
matched against them, is common especially in horticulture. In these cases
the number of classes is higher than is normally to be advocated for other
arbitrary class divisions. It may be found, however, that the range of variation
in population samples is so great as not to repay the labour involved, unless
colour differences appear to show some significant relationship with, say,
environmental factors . Other methods of colorometric analysis maybe
employed, such as measurement of amounts of pigment extracted per unit .
area, but as Heslop-Harrison (l952b) points out, the amount of pigment
present may not bear a linear relationship to the visual effect it produces. The
interpretation of colour will depend on what is known about the way in
which it is geneticaIly controlled, ontogenetically and environmentally
modified.
Indumentum can similarly be divided into arbitrary classes or compared
against certain standards. Here again problems of terminology arise if it is
divided into named classes such as pubescent, hirsute, sericeous, etc., since
they are often used imprecisely and are in faot to some extent overlapping and
interchangeable. Detailed analysis of indumentum often reveals differences
in hair length, shape, number of cells, presence or absence of glandular
trichomes, posture, etc. (see p. 154). .
Sculpturing (as in seeds and fruits) and patterning (as for example the
maculation in Digitalis purpurea corollas) are more difficult to assess. They
can, given time and patience, be specified by elaborate geometrical analysis.
The problems involved here are discussed by Heslop-Harrison (1952) and
exemplified by his series of studies on Dactylorchis populations (1948, 1951,
1952a, 1953a). . .
Division of numerically assessable characters into arbitrary classes may
also be a useful procedure. This method has the advantage that it is a great
deal more rapid than very precise scoring and 'is probably as valuable,
although care must be taken, through adequate sampling, to see that the
arbitrary grades are not absurd.
Quantitative characters appear to be more frequent iIi the lower taxonomic
categories, at specific and infra-specific levels. Thus species may be separated
by leaf size, corolla length, seed weight, etc., characters which are quantita
tively assessed and seldom of use at higher levels in the hierarchy._Most
morphological differences between ecotypes (and subspecies), for example,
hll.Ve been found to be of quantitative nature and may depend on polygenic
or multiple-gene inheritance; they may in fact only be appreciated after
DEFINITION OF CHARACTERS 119
statistical analysis. On the other hand, family characters tend to be
qualitative.
3. "Good" and "Bad" Characters
All characters arc theoretically of use in classification but experience has
shown that their value or use varies according to a number of factors and
according to the type of taxonomic activity envisaged. 'Great reliance is
placed on "good" characters, at least in practical taxonomy. Good characters
are those tl\at (1) are not subject to wide variation within the samples being
considered; (2) do not have a high intrinsic genetic variability; (3) are not
easily susceptible to environmental modification; (4) show consistency,
i.e. agree with the correlations of characters existing in a natural system 'of
classification which was constructed without their use.
We will consider this last point first. The consistency of a character
will help us to make a decision between two alternative classifications: it can
help us to resolve differences. In constructing a classification the taxonomist
is frequently faced with conflicting evidence. In these circumstances a
guiding principle would appear to beto select for the purpose ofgroup-making '
that character or group of characters which agrees with the overall facies of the
consequent groups. This is what we have referred to in Chapter 2 as correla- ,
lion or a posteriori weighting of characters,. It is essentially an intuitive or
neural use of Adansonian methods of makmg classes according to maximum
co-variation of attributes, except that facies is substituted for analysis of
attributes. Ther.:: are two aspects of this use of characters to be distinguished:
firstly, there is the value of the character as regards making a choice between '
possible alternative groupings, and secondly, the use of the character as a
diagnostic, i.e. for its predictive value. It is in the latter sense that it is more
useful to regard a character as reliable since its use will allow us to make a large
num'ber of deductions about the other attributes of the groups it off.
In b0th senses we are dealing with characters which have already been selected
for taxonomic use and which are being then considered for a particular
purpose.
It should be pointed out that there is a stroni element of circular argument
in this procedure of relying upon a character because it agrees with an existing
classification and then saying that the classification must be a good one, since
one may be doing no more than bolster up .the errors of that classification.
Ideally the whole classification should be broken down and the new character
induded along with the others for the purposes of completely estimating the
correlations anew. ' '
The other aspects of good characters, which we may loosely term their
fixity, is concerned with the selection of characters for general usc. Whether
for diagnosis or synthesis, a good charactcr must obviously be one that is
little influenced by environmental ' changes. The range of expression of a
is genetically determined and the actual expression will depend on
120 THE CONCEPT OF CHARACTERS [Ch.4
the environment-genotype reaction within that range. Good characters will
have ltllarrOW range of expression and will be easily recognizable. *
The inherent genetic variability of characters is a subject about which we
often have very inadequate information. As a general rule we would agree '
with Rollins (1958) that "we wish to rely on those characters as indicators of
relationship (or lack of it) that are so deeply seated in the genetic constitution
of the species that they cannot be easily obliterated or greatly modified by
the direct effects of any given, simply segregating factor or combination of
segregating factors". The effects of simple gene differences on morphological
features will be considered in more detail in Chapter 11. It IS sufficient to
mention ' here that many of the characters regarded by . taxonomists as
fundamental may be found to vary within populations and be inherited by
simple genetic mechanisms. A well-known example is described in Layia
spp. (Compositae subtrib. Madiinae) where 1- or 2-factorial inheritance
governs the presence or absence of ray flowers and paleae,
characters which are extensively treated by taxonomists as having tribal
significance (Clausen, 1951; Clausen & Hiesey, 1958a). Other features which
may be more important from a genetical and evolutionary viewpoint may
have been ignored by taxonomists.
It is, however, important to distinguish between those characters which
can and do vary by simple genetical mechanisms in a population and those
which are shown to be simply based genetically but which are not in fact
easily changed.
Thus the glabrous-fruited condition in Dithyrea wislezenii (Cruciferae),'
which was the main basis for describing D. griffithsii as a distinct species, is
a simply inherited character under single gene control. The pubescent plants
studied by Rollins (1958) were all homozygous and recessive for the pubescent
fruit character and he concluded from his studies that presence or absence of
pubescence on the fruits of D. wislezenii is of no taxonomic significance.
Here the character in question is no't only simply inherited but shows a
high likelihood of change. t
It would appea( to follow from this discussion that we should rely on
those characters that depend on a multiplicity of genes or gene combinations
for their ultimate expression, but great caution should be exercised in applying
this generalisation since it is not just the genetic basis of a character that has
to be assessed but its possibility of change in practice and the effects of such
changes on the continued integrity of taxa. In other words, a taxonomic
character is only as good as its constancy, no matter what its genetic basis.
Plastic characters may be used to differentiate species if their ranges do not
overlap or if the overlap is not too extensive.
t In other Crucifers glabrous and hairy-fruited variants can form pure popula
tions or' grow mixed together-presumably depending on whether the initial
coloniser was homozygous or heterozygous for the character in question (e.g. /saris,
Sisymbrium). The same often applies to albino variants in many families.
FUNCTION AND TAXONOMIC CHARACTERS 121
Other asptCts of genetic control of characters in assessing their value,
including pleiotropism causing mUltiple effects, are discussed in Chapter 11.
As we have already mentioned in Chapter 1, there-is a tendency to rely on
certain kinds of characters, such as those pertaining to the flower, in con
structing classifications. In terms of consistency and lack of plasticity floral
characters probably still justify their primary role, but this is not to deny that
other kinds of characters, notably vegetative ones, mlty not be equally
important in many cases. More fundamental considerations of floral features
are concerned with the concepts of constitutive and biological characters
which are discussed in the next section.
FUNCTION AND TAXONOMIC CHARACTERS
For many purposes today the plant taxonomist is seldom concerned with
the functional value of the characters he deals with in delimiting, describing
and classifying. In practice his concern is more with the constancy, reliability
and lack of variation or characters, as we have just seen, than with their
possible adaptive or functional significance (Heywood, 1959a). It was, however,
a characteristic of most pre-Darwinian taxonomy that the characters chosen
to construct classifi<;ations were intended to be those of high physiological or
functional value (the original "natural" classifications based on the true
nature or essences of organs, cf. Cain, 1958, 1959c). In fact, so little was known
about physiological importance that not much more than lip service could
be paid to these precepts. The gradual rejection of physiological characters
as having a function in taxonomy, by de Candolleand later taxonomists, has
been discussed in Chapter 1.
Some recent writers have pointed out that the role of function in taxonomy
has been unduly neglected (Stebbins, 1959b; Cain, 1959c), and Huxley has
introduced the concept of evolutionary grades in. an amplification of earlier
ideas on progressive evolution (anagenesis of Rensch) which bears on the same
problem. It was Wernham (1912), however, who introduced the distinction
between biological and fortuitous characters in his studies pri the evolutionary
origin of the sympetalous Dicotyledons. Biological cfiaracters are those
related to some vital function or advantage; they are usually floral, such
as epigyny, zygomorphy, sympetaly, etc. These biological characters were
those involved in progressive evolution, marked by two fundamental ten
dencies-economy in production of reproductive parts and adaptation to
insect visitors. Fortuitous characters, on the other hand, showed no such
(apparent) functional relationship.
Before considering the validity of this distinction, it is worth pointing out
that many Angiosperm families are recognised and described largely on the
Basis of such biological characters. This receives some explanation from the
observation by Stebbins (1950,1951) that the relative simplicity of plant struc
ture makes for a large amount of parallel variation, and families are made up,
as regards floral features, of combinations of the foUowing pairs of features:
122 T H ~ CONCEPT OF CHARACTERS [Ch.4
presence or absence of sepals and/or petals ; polypetaly versus sympetaly;
actinomorphy versus zygomorphy; numerous stamens versus few stamens;
apocarpy versus syncarpy; numerous Qvules versus few ovules; axile versus
parietal placentation; superior versus inferior ovary. Only 860f the possible
256 combinations of these characters were represented in the 438 families and
part-families studied. Nearly half of the total families (or part-families)
considered fell into one of twelve combinations indicating a strong deviation
from random representation. In other words, there is little scope for diver
sification with such limited themes and it is precisely such features that
Wernham had in mind when discussing biological characters. These selfsame
characters were those distinguished by Diels as constitutive and by other
authors as organisational characters; they are constant throughout a wide
range of individuals, species, genera, etc., that is, throughout a higher taxon.
Their taxonomic value is high and, as Sporne (1956) says, they are character
istic of a large area of ilJfinity. They are contrasted with non-constitutive
features which are less constant and are subject to environmental modification.
As we have noted, the number of "basic" characters in Angiosperm flowers
is limited and they evolved more than once in separate lines. Epigyny,
sympetaly, etc. may therefore be considered as adaptive convergences in the
Angiosperms. This has serious taxonomic implications, as van der Pijl has
pointed out. The occurrence of such features in flowers otherwise considered
unspecialised (e.g. epigynyor perigyny in Vict.orU:, C.alycanthus; syn:Petaly.in
some Annonaceae) should not be taken as ah mdlcatlOn of taxonomic affimty
and used as a motive for the displacement of taxa with such features to other
parts of the taxonomic system. In other words, such features should not be
weighted. It seems in fact probable that many Angiosperm families are
better regarded as grades as a result of parallel and convergent evolution.
Whether they are, as a result, unnatural is a question requiring further
research.
Although constitutive, organisational and biological characters are often
regarded as synonymous (cf. Spbrne, 1956) and the particular examples
chosen to illustrate them in fact coincide, they differ in the functional value
ascribed to them. As we have seen, Wernham's biological characters implied
a functional relationship with the environment, i.e. they were adaptive;
constitutive characters on the other hand are regarded (e.g. by Lam, 1959) as
non-adaptive. The explanation perhaps lies in the confusion that surrounds
the use of the term adaptation. An adaptation is generally defined as a
feature of an organism that confers some advantage on it, while adaptation
is the acquisition of such a feature by organisms. The words therefore refer
both to a process and to the result of that process; as far as thc organism is
concerned, adaptation is active or passive (Mayr, 1942)-it is happening
now or has happened in the past and is being exploited now. Function and
functional are used in essentially the same sense. As Simpson (1953a) notes, it is
a relationship between organism and environment that makes a charaCter
\ ~
FUNCTION 'AND TAXONOMIC CHARACTERS 123
advantageuus. It IS difficult to ' define adaptation more strictly without
finding that the definition hinges almost entirely on the degree to which we
can recognise whitt may be considered an ) ldaptive relationship. In other
words, while conspicuous relationship betwcen organism and environment
(as for example in prostrate maritime races of inland species, such as Cytisus
scoparius subsp.maritimus) present no problem, in the majority of cases the
acceptance of the adaptive nature of particular of,il plant depends
on the ingenuity, perception, skill and indeed imagination of the botanist
(cf. Barber, 1956).
For most of this century a negative and somewhat extremist attitude to
function and taxonomic characters has been adopted, largely for historkal
reasons (cf. Straw, 1956; van der Pijl, But it would appear to stem
more from ignorance than positive knowledge. Barber (1955) goes so far as
to say that "the question of the value of the morphological charac
teristics which the taxonomist uses to separate species populations is still
one of the crucial problems of evolutionary biology". He further comments,
with some that the museum or herbarium taxonomist often
regards diagnostic characters as having no selective value, if he considers the
problem at all. He is more concerned with the constancy of the features he has
chosen for discrimination between_groups (Heywood, 1959a). Quite apart from
the problems of whether such characters may be interpreted in terms of
function, some botanists doubt the value, for taxonomic separation, of
characters which can be shown to be adaptive (Borril1, 1961c). * From an
evolutionary viewpoint, Barber considers there are four possible explanations
concerning the nature of Jiagnostic characters:
(1) They are the result of genetic drift causing random fixation of genes
and may therefore have no adaptive value.
(2) They are the byproducts of the activity of other genes (pleiotropic
gene-action). Selection for one character may be due ' to the pleiotropic
effect of the ge'ne controlling another.
(3) They are the result of developm-ental correlations such as allometric
growth. If increase in size, for example, arjses through selection, any feature
correlated developmentally with size will also be subject to change; and if
these corresponding changes are selectively neutral they will persist.
(4) They have, or have had at some time during the history of the species,
some selective value and arose througlLnatur.al-"'Selection acting on random
mutations and recombinations.
A fifth possibility should be added. The diagnostic character may form
part of a complex of integrated adaptive characters. In other words, it may
belong to an adaptive complex based on an equilibrium due to the interaction
of many genes, often held together by linkage.
Very careful investigation is needed to determine which of the above
A viewpoint stemming from de Candolle's rejection of physiolbgical characters
for taxonomic comparison (cf. Cain, 1959a).
124 . THE CONCEPT OF CHARACTERS [Ch.4
mechanisms may have been involved. Adaptation shoUld be considered in
terms of ability to survive in a particular habitat and will therefore concern
such features as leaf size, internode developmerit, etc., features which are
greatly modified by the environment and therefore the least reliable. Adapta
tion depends, however, not only on survival of the individual buton the
propagation of the race. Reproductive structures will therefore be important
and the relative constancy of these structures may be promoted by natural
selection.
In the grass Melica mutica an apparently trivial taxonomic character-a
club-shaped rudiment at the apex'of the spikelets-hasa high selective value
as an effective dispersal mechanism since it is attractive to ants. Similarly in
the Hordeeae tribe of the Gramineae some of the most conspicuous of the
reproductive features used in the generic and specific taxonomy of the group
(shown by the glumes, lemmas and awns) consist of a series of different but
efficient seed-dispersal mechanisms. In the Compositae similar features with
similar selective values are used for taxonomic separation of genera. Stebbins
(1950) quotes examples from the tribe Cichorieae, where the floral structure of
the various genera is essentially similar and genera and species are diagnosed
by differences in the involucral bracts and in the mature fruits, especially in
the pappus of the cypselas ("achenes"). These are largely related to seed
protection and seed dispersal mechanisms: In the Chrysantheminae dimor
phism of the cypsela is related to different dispersal mechanisms, ,this feature
being one of the characters separating Chrysanthemum from related genera
sllch as Leucanthemum. Other examples are known in Hypochaeris, Crepis,
etc. in the same family. (See Salisbury, 1942, and Dansereau & Lems, 1957,
for a fuller consideration.)
In such groups as the Compositae and the Gramineae with reduced
flowers crowded into inflorescences, the whole inflorescence often evolves
as a unit adapted to the methods of pollination or dispersal, or both. Zohary
(1950) has outlined trends in the evolution of the Composite capitUlum in
which the involucre often opens like a capsule to release the cypselas (e.g.
Senecio vulgaris) or retains them and is itself the unit of dispersal (as in
Echinops and Centaurea squarrosa). In some cases (e.g. Crepis aspera) the
marginal cypselas are retained by the indurated involucral bracts while the
others are freely dispersed-thereby achieving different degrees of dispersal.
A similar result is obtained by the production of heteromorphic cypselas,
the outer being epappose and the inner pappose (Galinsoga parvij/ora), or the
pappus of the outer cypselas being caducous (Senecio jacobaea-cf. Burtt,
1961 a). L o ; ~ of pappus is not to be regarded as a retrograde loss of an efficient
mechanism but is often a specialisation. Although the pappus must have
played an important part in achieving the vast distribution of the Compositae,
under particular conditions it is disadvantageous (Burtt, 1961a).
One of the objections that has been raised to the hypothesis that elaborate
form and colour has evolved in response to pollinating agents is that many
125 FUNCTION AND TAXONOMIC CHARACTERS
such flowers are either selfing or apomictic. They almost certainly evolved,
however, from cross-pollinating ancestors-the change in breeding system
being brought about by changes in selective pressure.
Adaptive specialisation appears more clearly at the specific and parti
cularly infra-specific level: in populations showing adaptive response to
envirortmental conditions (Heywood, 1959a). This is discussed in detail in
Chapter 12. On the other hand, Simpson is of the opinion that the adaptive
significance of diagnostic characters becomes more apparent, at least in
animal groups, the higher the taxonomic category. Thus higher categories
should be distinguished by features of evident adaptive nature. This relation
ship is not, however, so obviously apparent in flowering plants, and zoologists
often express surprise that floral characters which, as we have mentioned,'
play such a marked role in higher-plant taxonomy have been seldom studied
from a functional or adaptational viewpoint.
As mentioned above, many people have denied the adaptive value of the
so-called biological (constitutive, organisational) characters, such as zygo
morphy, sympetaly, complex pollination mechanisms, etc. A survey of the
views of such anti-selectionists as Goebel, Troll, Good, and Nelsson is given
by van der Pijl (1960), who discusses their interpretation of the flower in terms
of "Gestalten", automatic autogenesis or physiological necessity. During the
last few decades there has been a renewal of research into floral biology,
particularly in tropical countries, using the tools of ecology, genetics, zoo
logical physiology, and it has been possible to demonstrate by experiment
that the intricacies of form, colour, odour and pattern in flowers have an
important role in attracting and orientating pollinating insects.
Van der Pijl points out that a major stumbling-block in considering the
flower from a functional point of view was that studies were made on
temperate floras where bees and butterflies were normal pollinators, and
where nectaries are considered a fundamental floral attraction. In other words
the climate was wrong both for understanding the flower and for interpreting
the problems. If we look at the earliest known fossil Angiosperms we find
that they show such features as we have previously been considering con
stitutive or organisational, e.g. epigyny, sympetaly, syncarpy in flowers
which are otherwise "unspecialised" or simple. It is difficult to see in what
way such features could be adaptive if we consider them in relation to their
present day pollinators. We now know, however, from palaeontological
research that in the period when the early were developing
(Lower Cretaceous and probably Upper Jurassic), beetles (Coleoptera) were
well differentiated and abundantly available as pollinators and that some
flies '(Diptera) were present. The Hymenoptera and Lepidoptera were not,
on the other hand, sufficiently well evolved to be of much significance as the
pollinators of early angiosperms. A good summary is given by Barnard (1961).
Parallelism of form in flowers known to be pollinated by similar agents
may in itself be regarded as evidence of an adaptive relationship. For example
126 THE CONCEPT OF CHARACTERS [Ch. 4
the flowers of Orchidaceae and Zingiberaceae, Polygalaceae and Pap ilion
oideae may be regarded as convergent adaptations in response to similar
methods of pollination. Bradshaw (1958) makes the same point about parallel
and repeated correlations between habitat and characters at the genecological
level as suggesting probable adaptation. On the other hand, characters which
are adaptive in a broad basic sense (e.g. the very features which define the
Angiosperms or the birds) are more likely to remain conservative for longer
than characters that have been evolved in relation to special conditions. For
example, those floral characters related to specialised pollinators (as in Salvia)
or to particular agents of dispersal (as in the subgenera of Trifolium) can be
fairly safely accepted as adaptive characters that have played little part in the
basic evolution of the higher groups and are associated more often with
generic or specific differentiation. The cushion habit and the evolution of
berries from capsules by the continued growth of the pericarp wall after fertili
sation are other characters than one can reasonably accept as "superficial"
specialisations that have had little to do with the basic evolution of families.
Each case must be judged on its own merits.
In summary, we may agree with Cain (1959c) that the taxonomic value of
characters should not be judged, for purposes of comparison, mainly in
terms of their function, following the early a priori taxonomists such as
Linnaeus and Caesalpinus; nor should we deny taxonomic value to characters
which can be demonstrated as functional and rely on those which have no
apparen,t functional value, as has been proposed by several more recent
botanists such as Sachs (1906), Wilmott (1950), and Borrill (196Ic).
As regards the constant characters of taxonomic groups, it is most
probable that this very constancy is maintained through the forces of natural
selection acting on the populations containing the characters, so that they
are more likely to have a selective value than not. That we cannot comprehend
in what way such characters may be functional does not mean they have
(or had) no adaptive significance.
For the purposes of phenetic comparisons, knowledge of the adaptive
nature of characters has little value except in so far as it stops us counting
the same character more than once through the functional correlation of
apparently separate characters. This question of the co-variation of characters
which are necessarily correlated is discussed below. The role of function in
the evolutionary interpretation of comparative data is, however, considerable.
CHARACTERS AND THE DEFINITION OF GROUPS
As mentioned above when referring to species, the characters chosen to
describe a taxonomic group form an abstract for pI actical convenience.
They do not in fact define the group. Similarly, as Simpson (l953a) says, the
diagnosis of a higher taxon (i.e. above the species level) is not a description
of any ancestor of the group. In other words, by abstracting from a higher
taxon those cnar&cters which are common to all its members, one may get
------- --- - - - - --- -
EVOLUTIONARY INTERPRETATION OF DATA 127
what is often referred to as a common ground-plan-some sort of general
basic pial). on which the group was modelled. This idealistic approach was
characteristic of certain schools of archetype seekers or typologists, parti
cularlythe Germanic school. But as Cain & Harrison (1960) point out, this
procedure may in fact give us less and less information about the presumed
ancestor of the group, since the rank of the group from which the characters
are abstracted is higher and contains a much greater diversity than any
ancestral taxon. It is also quite possible that the commoll ancestor did not
itself possess these characters in common. Features it did possess may have
been lost in the course of evolution; or the common ancestor may have had
the characters shown by an existing member of the group which have
survived by stasigenetic persistence. It is of course possible, too, that the
group may not have had a common ancestor at all if the resemblances are
partly the result of convergence (cf. Sporne, 1956).
Apart from considerations of common ancestors, it is by considering the
abstracted characters as in some way representing and defining the group
as a whole that we fall into serious typological error. A genus or higher
taxon is not defined by possessing a particular combination of floral and
vegetative features: it can be differentiated or recognised by these. These
characters are the ones which the component species share in common and
it is useful for the practical purposes of characterisation, identification and
keying that they be abstracted. (It should, however, be pointed out that since
phenetically determined groups are based on overall affinity, it is possible
that in some cases particular features may be modified or even lost in some
of the constituent members, with the result that the group cannot be diagnosed
(cf. Cain, 1954b; discussion in Simpson, 1961, pp. 93-103) but only described.
A number of currently accepted genera probably come under this heading.)
They are not, however, the only characters the group has in common, nor
indeed the only diagnostic characters since others may remain unrecognised
or be too difficult for practical use.
EVOLUTIONARY INTERPRETATION OF COMPARATIVE DATA
So far we have been concerned with the arrangement of comparative data
derived from all possible sources so as to produce a phenetic classification
based on overall similarity. The groups in this classification are recognised
by their resemblances without any weighting. This is an essential first step
in the production of an evolutionary or "phyletic" classification, although it
will not by itself produce one. Further interpretation of the phenetic features
of the groups and additional procedures are necessary if a phylogenetic
classification is aimed at. It may not be possible in some groups to go beyond
a phenetic arrangement, but usually it is possible to derive some phyletic
information from' the comparative data. The two most important kinds of
information that can be sought are those referring to convergence and
primitive characters. A further aspect to be considered is the necessary
128 THE CONCEPT OF CHARACTERS [Ch.4
correlation of characters as already mentioned; and the genetic evaluation
of characters and its bearing on the likelihood of their alteration or modifica
tion is also an increasingly fruitful source of information. Primitive characters
have already been discussed in Chapter 2 in detail and are not considered
further here.
1. Convergence
Convergent and parallel evolution has already been defined in Chapter 2.
The effects of convergence on the interpretation of natural classifications have
been underestimated in the past (Philipson, 1961) and, in flowering plants at
;east, the extent to which it has been unconsciously accepted in current
classifications is still not yet appreciated. If evolutionary divergence only
occurred in groups, then, as Cain (l959b) points out, a natural and phyletic
arrangement of it would virtually coincide. However, if convergence has
occurred, it might well result in forms being grouped together on the basis
of their convergent features although in respect of other features they would
be separated. In the absence of a fossil record it may be very difficult to tell
whether a natural arrangement is also phyletic, since the extent to which it
contains convergent forms cannot be determined. The assumption that
forms which are most like each other will be phyletically related in the sense
that they have a common ancestor is basically valid, but it does not allow
for the distortion of similarity by convergence which may not even be sus
pected in the group in question.
The Adansonian method of classification, when thoroughly applied, is
likely to overcome the effects of convergence in that the more thoroughly
the comparisons are made, using as many kinds of data as possible, the less
likely are convergent forms to affect the result. In other words, the large
numbers of different features used will probably compensate for any con
vergent similarities. Groups which on the basis of their general affinities are
widely separated will show resemblances in only the few convergent characters
By accepting these convergent characters as indicative of phyletic relationship
we would clearly be weighting them, and this is of course contrary to the
principles of general Adansonian classification. Unfortunately this weighting
procedure is nowadays frequently used, especially when newer kinds of
evidence are employed. As a result phenetic arrangements become distorted
and pseudo-phylogenetic classifications are proposed. The normal justification
of this procedure is that the new line of evidence (such as biochemical,
cytological, etc.) is regarded as more important, fundamental or at least
more trustworthY" than the traditional characters. There are two objections
to this procedure: firstly, as already mentioned, in phenetic comparisons
characters are not weighted, largely because we have no valid criteria for
jUdging the relative importance of different kinds of comparative data;
secondly, there is no reason to believe that these newer features are not also
susceptible to deviation and even reversal, so that reliance on